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Enzyme Name | UniProtKB | KEGG |
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| P07987 | Q9C1S9 |
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Protein name | Exoglucanase 2 | Exoglucanase-6A | cellulose 1,4-beta-cellobiosidaseexo-cellobiohydrolasebeta-1,4-glucan cellobiohydrolasebeta-1,4-glucan cellobiosylhydrolase1,4-beta-glucan cellobiosidaseexoglucanaseavicelaseCBH 1C1 cellulasecellobiohydrolase Icellobiohydrolaseexo-beta-1,4-glucan cellobiohydrolase1,4-beta-D-glucan cellobiohydrolasecellobiosidase |
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Synonyms | EC 3.2.1.91Exoglucanase IIExocellobiohydrolase IICBHII1,4-beta-cellobiohydrolase | EC 3.2.1.91Exocellobiohydrolase 6A1,4-beta-cellobiohydrolase 6ABeta-glucancellobiohydrolase 6AAvicelase 2 |
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Pfam | PF00734 (CBM_1) PF01341 (Glyco_hydro_6) [Graphical view]
| PF00734 (CBM_1) PF01341 (Glyco_hydro_6) [Graphical view]
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CAZy | GH6 (Glycoside Hydrolase Family)
| GH6 (Glycoside Hydrolase Family)
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KEGG pathways | MAP code | Pathways |
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MAP00500 | Starch and sucrose metabolism |
UniProtKB:Accession Number | P07987 | Q9C1S9 |
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Entry name | GUX2_TRIRE | GUX6_HUMIN |
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Activity | Hydrolysis of 1,4-beta-D-glucosidic linkages in cellulose and cellotetraose, releasing cellobiose from the non- reducing ends of the chains. | Hydrolysis of 1,4-beta-D-glucosidic linkages in cellulose and cellotetraose, releasing cellobiose from the non- reducing ends of the chains. |
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Subunit |
| Monomer. |
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Subcellular location | Secreted. |
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Cofactor |
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References for Catalytic Mechanism | References | Sections | No. of steps in catalysis |
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[14] | Scheme 1 | 1 | [15] | Fig.6, Fig.7, p.8888-8890 | 1 |
references | [1] |
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Comments | X-RAY CRYSTALLOGRAPHY (2.0 ANGSTROMS). |
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Medline ID | 90333255 |
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PubMed ID | 2377893 |
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Journal | Science |
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Year | 1990 |
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Volume | 249 |
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Pages | 380-6 |
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Authors | Rouvinen J, Bergfors T, Teeri T, Knowles JK, Jones TA |
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Title | Three-dimensional structure of cellobiohydrolase II from Trichoderma reesei. |
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Related UniProtKB | P07987 |
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[2] |
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PubMed ID | 1761039 |
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Journal | Eur J Biochem |
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Year | 1991 |
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Volume | 202 |
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Pages | 367-77 |
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Authors | Gilkes NR, Claeyssens M, Aebersold R, Henrissat B, Meinke A, Morrison HD, Kilburn DG, Warren RA, Miller RC Jr |
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Title | Structural and functional relationships in two families of beta-1,4-glycanases. |
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[3] |
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PubMed ID | 8399160 |
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Journal | Biochemistry |
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Year | 1993 |
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Volume | 32 |
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Pages | 9906-16 |
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Authors | Spezio M, Wilson DB, Karplus PA |
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Title | Crystal structure of the catalytic domain of a thermophilic endocellulase. |
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[4] |
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PubMed ID | 8192865 |
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Journal | Biotechnol Appl Biochem |
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Year | 1994 |
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Volume | 19 |
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Pages | 141-53 |
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Authors | Woodward J, Brown JP, Evans BR, Affholter KA |
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Title | Papain digestion of crude Trichoderma reesei cellulase: purification and properties of cellobiohydrolase I and II core proteins. |
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[5] |
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PubMed ID | 7733957 |
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Journal | Biochem Biophys Res Commun |
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Year | 1995 |
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Volume | 209 |
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Pages | 1046-52 |
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Authors | Shultz MD, Lassig JP, Gooch MG, Evans BR, Woodward J |
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Title | Palladium--a new inhibitor of cellulase activities. |
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[6] |
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PubMed ID | 7857933 |
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Journal | Biochemistry |
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Year | 1995 |
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Volume | 34 |
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Pages | 2220-4 |
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Authors | Damude HG, Withers SG, Kilburn DG, Miller RC Jr, Warren RA |
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Title | Site-directed mutation of the putative catalytic residues of endoglucanase CenA from Cellulomonas fimi. |
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[7] |
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PubMed ID | 8615816 |
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Journal | Biochem J |
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Year | 1996 |
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Volume | 315 |
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Pages | 467-72 |
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Authors | Damude HG, Ferro V, Withers SG, Warren RA |
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Title | Substrate specificity of endoglucanase A from Cellulomonas fimi: fundamental differences between endoglucanases and exoglucanases from family 6. |
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[8] |
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PubMed ID | 8856058 |
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Journal | Eur J Biochem |
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Year | 1996 |
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Volume | 240 |
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Pages | 584-91 |
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Authors | Harjunpaa V, Teleman A, Koivula A, Ruohonen L, Teeri TT, Teleman O, Drakenberg T |
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Title | Cello-oligosaccharide hydrolysis by cellobiohydrolase II from Trichoderma reesei. Association and rate constants derived from an analysis of progress curves. |
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[9] |
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PubMed ID | 9010760 |
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Journal | J Biochem (Tokyo) |
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Year | 1996 |
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Volume | 120 |
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Pages | 1123-9 |
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Authors | Amano Y, Shiroishi M, Nisizawa K, Hoshino E, Kanda T |
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Title | Fine substrate specificities of four exo-type cellulases produced by Aspergillus niger, Trichoderma reesei, and Irpex lacteus on (1-->3), (1-->4)-beta-D-glucans and xyloglucan. |
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[10] |
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Comments | X-RAY CRYSTALLOGRAPHY (2.0 ANGSTROMS). |
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Medline ID | 97029636 |
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PubMed ID | 8875646 |
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Journal | Protein Eng |
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Year | 1996 |
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Volume | 9 |
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Pages | 691-9 |
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Authors | Koivula A, Reinikainen T, Ruohonen L, Valkeajarvi A, Claeyssens M, Teleman O, Kleywegt GJ, Szardenings M, Rouvinen J, Jones TA, Teeri TT |
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Title | The active site of Trichoderma reesei cellobiohydrolase II: the role of tyrosine 169. |
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Related PDB | 1cb2 |
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Related UniProtKB | P07987 |
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[11] |
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PubMed ID | 8959766 |
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Journal | Protein Expr Purif |
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Year | 1996 |
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Volume | 8 |
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Pages | 399-400 |
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Authors | Koivula A, Lappalainen A, Virtanen S, Mantyla AL, Suominen P, Teeri TT |
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Title | Immunoaffinity chromatographic purification of cellobiohydrolase II mutants from recombinant trichoderma reesei strains devoid of major endoglucanase genes. |
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[12] |
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PubMed ID | 9204518 |
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Journal | Appl Biochem Biotechnol |
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Year | 1997 |
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Volume | 66 |
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Pages | 39-56 |
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Authors | Medve J, Stahlberg J, Tjerneld F |
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Title | Isotherms for adsorption of cellobiohydrolase I and II from Trichoderma reesei on microcrystalline cellulose. |
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[13] |
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Comments | X-ray crystallography |
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PubMed ID | 9662445 |
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Journal | FEBS Lett |
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Year | 1998 |
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Volume | 429 |
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Pages | 341-6 |
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Authors | Koivula A, Kinnari T, Harjunpaa V, Ruohonen L, Teleman A, Drakenberg T, Rouvinen J, Jones TA, Teeri TT |
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Title | Tryptophan 272: an essential determinant of crystalline cellulose degradation by Trichoderma reesei cellobiohydrolase Cel6A. |
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Related PDB | 1bvw |
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[14] |
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PubMed ID | 9882628 |
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Journal | Biochem J |
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Year | 1999 |
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Volume | 337 |
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Pages | 297-304 |
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Authors | Varrot A, Hastrup S, Schulein M, Davies GJ |
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Title | Crystal structure of the catalytic core domain of the family 6 cellobiohydrolase II, Cel6A, from Humicola insolens, at 1.92 A resolution. |
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[15] |
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Comments | X-ray crystallography |
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PubMed ID | 10413461 |
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Journal | Biochemistry |
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Year | 1999 |
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Volume | 38 |
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Pages | 8884-91 |
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Authors | Varrot A, Schulein M, Davies GJ |
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Title | Structural changes of the active site tunnel of Humicola insolens cellobiohydrolase, Cel6A, upon oligosaccharide binding. |
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Related PDB | 2bvw |
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[16] |
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PubMed ID | 10411622 |
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Journal | Eur J Biochem |
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Year | 1999 |
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Volume | 262 |
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Pages | 637-43 |
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Authors | Carrard G, Linder M |
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Title | Widely different off rates of two closely related cellulose-binding domains from Trichoderma reesei. |
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[17] |
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Comments | X-ray crystallography |
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PubMed ID | 10508787 |
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Journal | Structure Fold Des |
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Year | 1999 |
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Volume | 7 |
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Pages | 1035-45 |
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Authors | Zou J, Kleywegt GJ, Stahlberg J, Driguez H, Nerinckx W, Claeyssens M, Koivula A, Teeri TT, Jones TA |
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Title | Crystallographic evidence for substrate ring distortion and protein conformational changes during catalysis in cellobiohydrolase Ce16A from trichoderma reesei. |
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Related PDB | 1qk0,1qk2,1qjw |
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[18] |
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PubMed ID | 10794732 |
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Journal | Biochem J |
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Year | 2000 |
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Volume | 348 Pt 1 |
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Pages | 201-7 |
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Authors | Davies GJ, Brzozowski AM, Dauter M, Varrot A, Schulein M |
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Title | Structure and function of Humicola insolens family 6 cellulases: structure of the endoglucanase, Cel6B, at 1.6 A resolution. |
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[19] |
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PubMed ID | 10601873 |
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Journal | Eur J Biochem |
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Year | 2000 |
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Volume | 267 |
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Pages | 244-52 |
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Authors | Zhang S, Barr BK, Wilson DB |
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Title | Effects of noncatalytic residue mutations on substrate specificity and ligand binding of Thermobifida fusca endocellulase cel6A. |
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[20] |
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PubMed ID | 11179652 |
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Journal | FEMS Microbiol Lett |
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Year | 2001 |
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Volume | 195 |
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Pages | 197-204 |
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Authors | Lehtio J, Wernerus H, Samuelson P, Teeri TT, Stahl S |
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Title | Directed immobilization of recombinant staphylococci on cotton fibers by functional display of a fungal cellulose-binding domain. |
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[21] |
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PubMed ID | 11325554 |
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Journal | FEMS Microbiol Lett |
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Year | 2001 |
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Volume | 198 |
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Pages | 57-63 |
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Authors | Limon MC, Margolles-Clark E, Benitez T, Penttila M |
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Title | Addition of substrate-binding domains increases substrate-binding capacity and specific activity of a chitinase from Trichoderma harzianum. |
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[22] |
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PubMed ID | 12007616 |
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Journal | Biochim Biophys Acta |
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Year | 2002 |
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Volume | 1596 |
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Pages | 366-80 |
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Authors | Tuohy MG, Walsh DJ, Murray PG, Claeyssens M, Cuffe MM, Savage AV, Coughlan MP |
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Title | Kinetic parameters and mode of action of the cellobiohydrolases produced by Talaromyces emersonii. |
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[23] |
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PubMed ID | 12188666 |
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Journal | J Am Chem Soc |
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Year | 2002 |
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Volume | 124 |
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Pages | 10015-24 |
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Authors | Koivula A, Ruohonen L, Wohlfahrt G, Reinikainen T, Teeri TT, Piens K, Claeyssens M, Weber M, Vasella A, Becker D, Sinnott ML, Zou JY, Kleywegt GJ, Szardenings M, Stahlberg J, Jones TA |
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Title | The active site of cellobiohydrolase Cel6A from Trichoderma reesei: the roles of aspartic acids D221 and D175. |
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[24] |
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PubMed ID | 12565856 |
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Journal | Biochem Biophys Res Commun |
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Year | 2003 |
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Volume | 301 |
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Pages | 280-6 |
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Authors | Murray PG, Collins CM, Grassick A, Tuohy MG |
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Title | Molecular cloning, transcriptional, and expression analysis of the first cellulase gene (cbh2), encoding cellobiohydrolase II, from the moderately thermophilic fungus Talaromyces emersonii and structure prediction of the gene product. |
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comments | This family belongs to Glycosidase family-6, which has an inverting mechanism (equatorial to axial conformation). Furthermore, this enzyme belongs to exoglucanase, which requrires polymer-chain ends for catalytic activity, whilst its family member (EC 3.2.1.4) is endoglucanase. The literature [7] suggests that endoglucanases from this family-6 may function by different mechanisms from this exoglucanase from the same family. The literature [15] confirmed that Asp226 (2bvw) atcs as proton donor (acid catalyst) and Asp405 as the catalytic base. In sharp contrast, the literature [6] doubted the role of Asp405, which suggests Asp268 might act as base catalyst. In any case, the identification of catalytic base is difficult for these enzymes.
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created | updated |
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2002-10-15 | 2009-09-29 |
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